RESEARCH

POPULATION GENETICS fromtheanalysisdatasetsindividualscoveredby <10,000 SNPs, with evidence of contamination, or first-degree relatives of others (table S1). We analyzed the data with principal components The genomic history of the Iberian analysis (PCA) (Fig. 1, C and D), f-statistics (11), and qpAdm (12) and summarize the results in Peninsula over the past 8000 years Fig. 1E. We confirmed the robustness of key findings by repeating analyses after remov- Iñigo Olalde1*, Swapan Mallick1,2,3, Nick Patterson2, Nadin Rohland1, ing SNPs in CpG dinucleotides (table S5) that Vanessa Villalba-Mouco4,5, Marina Silva6, Katharina Dulias6, Ceiridwen J. Edwards6, are susceptible to cytosine-to-thymine errors Francesca Gandini6, Maria Pala6, Pedro Soares7, Manuel Ferrando-Bernal8, even in UDG-treated libraries (1). Nicole Adamski1,3, Nasreen Broomandkhoshbacht1,3, Olivia Cheronet9, Previous knowledge of the genetic structure Brendan J. Culleton10, Daniel Fernandes9,11, Ann Marie Lawson1,3, of Mesolithic Iberia comes from three individ- 1,2,3 1,3 1,3 1 2 Matthew Mah , Jonas Oppenheimer , Kristin Stewardson , Zhao Zhang , uals from the northwest: LaBraña1 ( ), Canes1 5 5 Juan Manuel Jiménez Arenas12,13,14, Isidro Jorge Toro Moyano15, ( ), and Chan ( ). We add LaBraña2, who was a Domingo C. Salazar-García16, Pere Castanyer17, Marta Santos17, Joaquim Tremoleda17, brother of the previously reported LaBraña1 (figs. S1 and S2 and table S6), as well as Cueva Marina Lozano18,19, Pablo García Borja20, Javier Fernández-Eraso21, de la Carigüela (fig. S10), Cingle del Mas Nou, José Antonio Mujika-Alustiza21, Cecilio Barroso22, Francisco J. Bermúdez22, and Cueva de la Cocina from the southeast. In Enrique Viguera Mínguez23, Josep Burch24, Neus Coromina24, David Vivó24, northwest Iberia, we document a previously un- Artur Cebrià25, Josep Maria Fullola25, Oreto García-Puchol26, Juan Ignacio Morales25, appreciated ancestry shift before the arrival of F. Xavier Oms25, Tona Majó27, Josep Maria Vergès18,19, Antònia Díaz-Carvajal28, Downloaded from 28 25 29,30,31 farming (Fig. 2A, fig. S5, and table S7). The oldest Imma Ollich-Castanyer , F. Javier López-Cachero , Ana Maria Silva , individual Chan was similar to the ~19,000-year-old 32 33 32 Carmen Alonso-Fernández , Germán Delibes de Castro , Javier Jiménez Echevarría , El Mirón, whereas the La Braña brothers from 34,35 36 Adolfo Moreno-Márquez , Guillermo Pascual Berlanga†, Pablo Ramos-García , ~1300 years later were closer to central European 34 34 37 José Ramos-Muñoz , Eduardo Vijande Vila , Gustau Aguilella Arzo , hunter-gatherers like the Hungarian KO1, with Ángel Esparza Arroyo38, Katina T. Lillios39, Jennifer Mack40, Javier Velasco-Vázquez41, an even more extreme shift ~700 years later in 42 43,44 45 Anna Waterman , Luis Benítez de Lugo Enrich , María Benito Sánchez , Canes1. This likely reflects gene flow affecting http://science.sciencemag.org/ Bibiana Agustí46,47, Ferran Codina47, Gabriel de Prado47, Almudena Estalrrich48, northwest Iberia but not the southeast, where Álvaro Fernández Flores49, Clive Finlayson50,51,52,53, Geraldine Finlayson50,52,53, individuals remained close to El Mirón (Fig. 2A). Stewart Finlayson50,54, Francisco Giles-Guzmán50, Antonio Rosas55, More data from the Mesolithic period, especially Virginia Barciela González56,57, Gabriel García Atiénzar56,57, Mauro S. Hernández Pérez56,57, from currently unsampled areas, would provide Armando Llanos58, Yolanda Carrión Marco59, Isabel Collado Beneyto60, additional insight into the geographical impact David López-Serrano61, Mario Sanz Tormo†, António C. Valera62, Concepción Blasco43, and archaeological correlates of this ancestry shift. Corina Liesau43, Patricia Ríos43, Joan Daura25, María Jesús de Pedro Michó63, For the Neolithic and Copper Age, we model Agustín A. Diez-Castillo64, Raúl Flores Fernández†, Joan Francès Farré65, populationsasmixturesofgroupsrelatedto Rafael Garrido-Pena43, Victor S. Gonçalves30, Elisa Guerra-Doce33, Anatolian Neolithic, El Mirón, and KO1 (Fig. 2A 66 67 68 and table S8). We replicate previous findings of

Ana Mercedes Herrero-Corral , Joaquim Juan-Cabanilles , Daniel López-Reyes , the arrival of Anatolian Neolithic–associated an- on March 14, 2019 Sarah B. McClure69, Marta Merino Pérez70, Arturo Oliver Foix37, cestry in multiple regions of Iberia in the Early Montserrat Sanz Borràs25, Ana Catarina Sousa30, Julio Manuel Vidal Encinas71, Neolithic (7, 8, 12); however, sampling from this Douglas J. Kennett10,69, Martin B. Richards6, Kurt Werner Alt72,73, 4,74 9 8 1,2,3 period remains limited and studies of larger sam- Wolfgang Haak , Ron Pinhasi , Carles Lalueza-Fox *, David Reich * ple sizes and additional sites will be important to shed further light on the interaction between We assembled genome-wide data from 271 ancient Iberians, of whom 176 are from the the incoming farmers and indigenous hunter- largely unsampled period after 2000 BCE, thereby providing a high-resolution time transect gatherers. For the Middle Neolithic and Copper of the Iberian Peninsula. We document high genetic substructure between northwestern Age, we reproduce previous reports of an in- and southeastern hunter-gatherers before the spread of farming. We reveal sporadic crease of hunter-gatherer–related ancestry after contacts between Iberia and North Africa by ~2500 BCE and, by ~2000 BCE, the 4000 BCE (6, 7, 12, 13), with higher proportions ’ replacement of 40% of Iberia s ancestry and nearly 100% of its Y-chromosomes by people in groups from the north and center. Using our with Steppe ancestry. We show that, in the Iron Age, Steppe ancestry had spread not only observations about population substructure in – – into Indo-European speaking regions but also into non-Indo-European speaking ones, and theMesolithicasareferenceframe,weshowthat we reveal that present-day Basques are best described as a typical Iron Age population the hunter-gatherer–related ancestry during those without the admixture events that later affected the rest of Iberia. Additionally, we document periods was more closely related to later north- how, beginning at least in the Roman period, the ancestry of the peninsula was transformed western (Canes1-like) hunter-gatherers than to by gene flow from North Africa and the eastern Mediterranean. the El Mirón–like hunter-gatherers (Fig. 2A), pro- viding clues about the source of this ancestry. he Iberian Peninsula, lying at the extreme single-nucleotide polymorphisms (SNPs) (2, 3) Our Copper Age dataset includes a newly re- southwestern corner of Europe, provides to generate genome-wide data from 4 Mesolithic, ported male (I4246) from Camino de las Yeseras an excellent context in which to assess the 44 Neolithic, 47 Copper Age, 53 Bronze Age, (14) in central Iberia, radiocarbon dated to T final impact of population movements en- 24 Iron Age, and 99 historical-period Iberians 2473–2030 calibrated years BCE, who clusters tering the continent from the east as well (Fig.1,AandB,andtablesS1andS2).Wealso with modern and ancient North Africans in the as interactions with North Africa. To study the generated 26 radiocarbon dates (table S3). We PCA (Fig. 1C and fig. S3) and, like ~3000 BCE genetic impact of prehistoric and historic events co-analyzed the new genomic data with previ- Moroccans (8), can be well modeled as having in Iberia, we prepared next-generation sequenc- ously reported data from 1107 ancient individ- ancestry from both Late Pleistocene North Africans ing libraries treated with uracil-DNA glycosylase uals, including 132 from Iberia (Fig. 1B) (2, 4–9), (15)andEarlyNeolithicEuropeans(tablesS9and (UDG) (1) and enriched them for ~1.2 million and 2862 present-day individuals (10). We filtered S10). His genome-wide ancestry and uniparental

Olalde et al., Science 363, 1230–1234 (2019) 15 March 2019 1of5 RESEARCH | REPORT markers (tables S1 and S4) are unique among cestry on the X-chromosome (table S14 and fig. in Western Europe but overlap genetically with Copper Age Iberians, including individuals from S7), a paradigm that can be exemplified by a Iron Age populations (Fig. 1D) showing substan- sites with many analyzed individuals such as Bronze Age tomb from Castillejo del Bonete tial levels of Steppe ancestry. Sima del Ángel, and point to a North African containing a male with Steppe ancestry and a In the historical period, our transect begins origin. Our genetic evidence of sporadic contacts femalewithancestrysimilartoCopperAge with 24 individuals from the 5th century BCE with North Africa during the Copper Age fits Iberians. Although ancient DNA can document to the 6th century CE from the Greek colony of with the presence of African ivory at Iberian that sex-biased admixture occurred, archaeolog- Empúries in the northeast (19)whofallinto sites (16) and is further supported by a Bronze ical and anthropological research will be needed two main ancestry groups (Fig. 1, C and D, and Age individual (I7162) from Loma del Puerco to understand the processes that generated it. fig. S8): one similar to Bronze Age individuals in southern Iberia who had 25% ancestry re- For the Iron Age, we document a consistent from the Aegean, and the other similar to Iron lated to individuals like I4246 (Fig. 1D and trend of increased ancestry related to Northern AgeIberianssuchasthosefromthenearbynon- table S16). However, these early movements and Central European populations with respect Greek site of Ullastret, confirming historical from North Africa had a limited impact on to the preceding Bronze Age (Figs. 1, C and D, sources indicating that this town was inhabited Copper and Bronze Age Iberians, as North and 2B). The increase was 10 to 19% (95% con- by a multiethnic population (19). The impact of African ancestry only became widespread in fidence intervals given here and in the percent- mobility from the central/eastern Mediterra- the past ~2000 years. ages that follow) in 15 individuals along the nean during the Classical period is also evident FromtheBronzeAge(~2200–900 BCE), we Mediterranean coast where non-Indo-European in 10 individuals from the 7th to 8th century CE increase the available dataset (6, 7, 17) from 7 to Iberian languages were spoken; 11 to 31% in two site of L'Esquerda in the northeast, who show 60 individuals and show how ancestry from the individuals at the Tartessian site of La Angorrilla a shift from the Iron Age population in the Pontic-Caspian steppe (Steppe ancestry) appeared in the southwest with uncertain language attri- direction of present-day Italians and Greeks (Fig. throughout Iberia in this period (Fig. 1, C and D), bution; and 28 to 43% in three individuals at 1D) that accounts for approximately one-quarter Downloaded from albeit with less impact in the south (table S13). La Hoya in the north where Indo-European of their ancestry (Fig. 2C and table S17). The same The earliest evidence is in 14 individuals dated to Celtiberian languages were likely spoken (fig. S6 shift is also observed in present-day Iberians ~2500–2000 BCE who coexisted with local people and tables S11 and S12). This trend documents outside the Basque area and is plausibly a without Steppe ancestry (Fig. 2B). These groups gene flow into Iberia during the Late Bronze consequence of the Roman presence in the pe- lived in close proximity and admixed to form Age or Early Iron Age, possibly associated with ninsula, which had a profound cultural impact the Bronze Age population after 2000 BCE with the introduction of the Urnfield tradition (18). and, according to our data, a substantial genetic

~40% ancestry from incoming groups (Fig. 2B Unlike in Central or Northern Europe, where impact too. http://science.sciencemag.org/ and fig. S6). Y-chromosome turnover was even Steppe ancestry likely marked the introduction In contrast to the demographic changes in the more pronounced (Fig. 2B), as the lineages com- of Indo-European languages (12), our results Classical period, movements into Iberia during the mon in Copper Age Iberia (I2, G2, and H) were indicate that, in Iberia, increases in Steppe an- decline of the Roman Empire had less long-term almost completely replaced by one lineage, R1b- cestry were not always accompanied by switches demographic impact. Nevertheless, individual M269.Thesepatternspointtoahighercon- to Indo-European languages. This is consistent sites—for example, the 6th century site of Pla de tribution of incoming males than females, also with the genetic profile of present-day Basques l'Horta in the northeast—bear witness to events supported by a lower proportion of nonlocal an- whospeaktheonlynon-Indo-Europeanlanguage in this period. These individuals, archaeologically

1Department of Genetics, Harvard Medical School, Boston, MA, USA. 2Broad Institute of MIT and Harvard, Cambridge, MA, USA. 3Howard Hughes Medical Institute, Harvard Medical School,

Boston, MA, USA. 4Max Planck Institute for the Science of Human History, Jena, Germany. 5Departamento de Ciencias de la Antigüedad, Grupo Primeros Pobladores del Valle del Ebro on March 14, 2019 (PPVE), Instituto de Investigación en Ciencias Ambientales (IUCA), Universidad de Zaragoza, Zaragoza, Spain. 6Department of Biological and Geographical Sciences, School of Applied Sciences, University of Huddersfield, Huddersfield, UK. 7Centre of Molecular and Environmental Biology, Department of Biology, University of Minho, Braga, Portugal. 8Institute of Evolutionary Biology, CSIC-Universitat Pompeu Fabra, Barcelona, Spain. 9Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria. 10Department of Anthropology and Institutes of Energy and the Environment, The Pennsylvania State University, University Park, PA, USA. 11Research Center for Anthropology and Health, Department of Life Science, University of Coimbra, Coimbra, Portugal. 12Departamento de Prehistoria y Arqueología, Universidad de Granada, Granada, Spain. 13Instituto Universitario de la Paz y los Conflictos, Universidad de Granada, Granada, Spain. 14Department of Anthropology - Anthropologisches Institut and Museum, Universität Zürich, Zürich, Switzerland. 15Museo Arqueológico y Etnológico de Granada, Granada, Spain. 16Departamento de Geografía, Prehistoria y Arqueología, Grupo de Investigación en Prehistoria, (UPV-EHU)/IKERBASQUE-Basque Foundation for Science, Vitoria, Spain. 17Museu d'Arqueologia de Catalunya- Empúries, L'Escala, Spain. 18Institut Català de Paleoecologia Humana i Evolució Social (IPHES), Tarragona, Spain. 19Àrea de Prehistòria, Universitat Rovira i Virgili (URV), Tarragona, Spain. 20Departamento de Prehistoria e Historia Antigua, Universidad Nacional de Educación a Distancia, Valencia, Spain. 21Departamento de Geografía, Prehistoria y Arqueología, Universidad del País Vasco, Vitoria, Spain. 22Fundación Instituto de Investigación de Prehistoria y Evolución Humana (FIPEH), Lucena, Spain. 23Área de Genética, Facultad de Ciencias, Universidad de Málaga, Málaga, Spain. 24Institut de Recerca Històrica, Universitat de Girona, Girona, Spain. 25SERP, Departament d’Història i Arqueologia, Facultat de Geografia i Història, Universitat de Barcelona, Barcelona, Spain. 26PREMEDOC Research Group, Departament de Prehistòria, Arqueologia i Historia Antiga, Universitat de València, València, Spain. 27Archaeom. Departament de Prehistòria, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain. 28Universitat de Barcelona-GRAMP/Museu Arqueològic de l'Esquerda, Roda de Ter, Spain. 29Laboratory of Prehistory, Research Center for Anthropology and Health, Department of Life Sciences, University of Coimbra, Coimbra, Portugal. 30UNIARQ, Faculdade de Letras, Universidade de Lisboa, Lisboa, Portugal. 31CEF, Department of Life Sciences, University of Coimbra, Coimbra, Portugal. 32Cronos S.C. Arqueología y Patrimonio, Burgos, Spain. 33Departamento de Prehistoria, Facultad de Filosofía y Letras, Universidad de Valladolid, Valladolid, Spain. 34Departamento de Historia, Geografía y Filosofía, Universidad de Cádiz, Cádiz, Spain. 35Departamento de Geografía, Historia y Humanidades, Universidad de Almería, Almería, Spain. 36School of Dentistry, University of Granada, Granada, Spain. 37Servicio de Investigaciones Arqueológicas y Prehistóricas de la Diputación de Castellón, Castelló de la Plana, Spain. 38GIR PrehUSAL, Departamento de Prehistoria, Historia Antigua y Arqueología, Universidad de Salamanca, Salamanca, Spain. 39Department of Anthropology, University of Iowa, Iowa City, IA, USA. 40Office of the State Archaeologist, University of Iowa, Iowa City, IA, USA. 41Departamento de Ciencias Históricas, Universidad de Las Palmas de Gran Canaria, Las Palmas, Spain. 42Mt. Mercy University, Cedar Rapids, IA, USA. 43Departamento de Prehistoria y Arqueología, Universidad Autónoma de Madrid, Madrid, Spain. 44Departamento de Prehistoria y Arqueología, Universidad Nacional de Educación a Distancia, Madrid, Spain. 45Departamento de Medicina Legal, Psiquiatría y Anatomía Patológica, Universidad Complutense de Madrid, Madrid, Spain. 46INSITU S.C.P., Centelles, Spain. 47Museu d'Arqueologia de Catalunya-Ullastret, Ullastret, Spain. 48Instituto Internacional de Investigaciones Prehistóricas de Cantabria IIIPC (Universidad de Cantabria-Gobierno de Cantabria-Santander), Santander, Spain. 49Arqueología y Gestión S.L.L., Fuentes de Andalucia, Spain. 50The Gibraltar National Museum, Gibraltar. 51Department of Anthropology, University of Toronto, Toronto, ON, Canada. 52School of Natural Sciences and Psychology, Liverpool John Moores University, Liverpool, UK. 53Institute of Life and Earth Sciences, University of Gibraltar, Gibraltar. 54Department of Life Sciences, Anglia Ruskin University, Cambridge, UK. 55Paleoanthropology Group, Department of Paleobiology, Museo Nacional de Ciencias Naturales (MNCN)–Consejo Superior de Investigaciones Científicas (CSIC), Madrid, Spain. 56Departamento de Prehistoria, Arqueología e Historia Antigua, Facultad de Filosofía y Letras, Universidad de Alicante, San Vicente del Raspeig, Spain. 57Instituto Universitario de Investigación en Arqueología y Patrimonio Histórico (INAPH), San Vicente del Raspeig, Spain. 58Instituto Alavés de Arqueología, Vitoria-Gasteiz, Spain. 59Departament de Prehistòria, Arqueologia i Historia Antiga, Universitat de València, València, Spain. 60Museu Arqueológic Vicent Casanova, Bocairent, Spain. 61Estrats, Treballs d'Arqueologia SL, El Campello, Spain. 62Era – Arqueologia, Oeiras, Portugal. 63Museu de Prehistòria de València, València, Spain. 64GRAM Research Group, Departament de Prehistòria, Arqueologia i Historia Antiga, Universitat de València, València, Spain. 65Museu i Poblat Ibèric de Ca n'Oliver, Cerdanyola del Vallès, Spain. 66Departamento de Prehistoria, Universidad Complutense de Madrid, Madrid, Spain. 67Museu de Prehistoria/SIP, Diputació de València, València, Spain. 68Arqueovitis sccl., Avinyonet del Penedès, Spain. 69Department of Anthropology, University of California, Santa Barbara, CA, USA. 70Unitat d'Antropologia Física, Departament de Biologia Animal, Facultat de Biologia, Universitat de Barcelona, Barcelona, Spain. 71Junta de Castilla y León, Servicio de Cultura de León, León, Spain. 72Center of Natural and Cultural Human History, Danube Private University, Krems, Austria. 73Department of Biomedical Engineering and Integrative Prehistory and Archaeological Science, Basel University, Basel, Switzerland. 74School of Biological Sciences, University of Adelaide, Adelaide, Australia. *Corresponding author. Email: [email protected] (I.O.); [email protected] (C.L.-F.); [email protected] (D.R.) †Independent researcher.

Olalde et al., Science 363, 1230–1234 (2019) 15 March 2019 2of5 RESEARCH | REPORT Downloaded from http://science.sciencemag.org/

on March 14, 2019

Fig. 1. Overview of the ancient Iberian genetic time transect. (A) Eurasian individuals (gray dots), with ancient individuals from Iberia and Geographic distribution and (B) dates of new and previously reported other regions (pale yellow) projected onto the first two principal samples. Random jitter is added for sites with multiple individuals. Sites components. (D) Section of the PCA in (C) marked with the dashed box. mentioned in the text are labeled. (C) PCA of 989 present-day west (E) Schematic representation of events documented in this study. interpreted as Visigoths, are shifted from those ancestry, probably related to the well-known evident in northeast Iberia in seven individu- at L'Esquerda in the direction of Northern and mobility patterns during the Roman Empire als from Sant Julià de Ramis from the 8th to Central Europe (Figs. 1D and 2C and table S18), (22) or to the earlier Phoenician-Punic pres- 12th centuries CE who, unlike previous ancient and we observe the Asian mitochondrial DNA ence (23);thelatterisalsosupportedbythe individuals from the same region, show North (mtDNA) haplogroup C4a1a also found in Early observation of the Phoenician-associated Y- African–related ancestry (Fig. 2C and table S19) Medieval Bavaria (20), supporting a recent link chromosome J2 (24). Gene flow from North and a complete overlap in PCA with present-day to groups with ancestry originally derived from Africa continued into the Muslim period, as Iberians (Fig. 1D). Central and Eastern Europe. is clear from Muslim burials with elevated North Our time transect allowed us to track frequen- In the southeast, we recovered genomic data African and sub-Saharan African ancestry (Fig. cy changes of phenotypically important variants from 45 individuals dated between the 3rd and 2D, fig. S4, and table S22) and from uniparental over the past 4000 years (fig. S9), a period that 16th centuries CE. All analyzed individuals fell markers typical of North Africa not present has been minimally sampled in the ancient DNA outside the genetic variation of preceding Iberian among pre-Islamic individuals (Fig. 2D and literature not just in Iberia but in Europe more Iron Age populations (Fig. 1, C and D, and fig. fig. S11). Present-day populations from south- generally. Before this work, it was known that S3) and harbored ancestry from both Southern ern Iberia harbor less North African ancestry the lactase persistence allele at rs4988235, which European and North African populations (Fig. (25) than the ancient Muslim burials, plausi- is present at moderate or high frequencies in 2D), as well as additional Levantine-related bly reflecting expulsion of moriscos (former most European populations today and is one of ancestry that could potentially reflect ancestry Muslims converted to Christianity) and repo- the strongest known signals of selection in from Jewish groups (21). These results demon- pulation from the north, as supported by histor- Europeans (26), occurred at extremely low fre- strate that by the Roman period, southern Iberia ical sources and genetic analysis of present-day quencies in Europe through the Bronze Age (2), had experienced a major influx of North African groups (25). The impact of Muslim rule is also raising the question of when it became common.

Olalde et al., Science 363, 1230–1234 (2019) 15 March 2019 3of5 RESEARCH | REPORT Downloaded from http://science.sciencemag.org/

on March 14, 2019

Fig. 2. Genome-wide admixture proportions using qpAdm. (A) Modeling (table S4). (C) Ancestry proportions for individuals from three sites in northeast Mesolithic, Neolithic, and Copper Age populations as a mixture of Anatolian Iberia dated between the 6th and 12th centuries CE. n represents the number of Neolithic, El Mirón, and KO1. Percentages indicate proportion of El Mirón + individuals analyzed in each site. (D) Ancestry proportions for individuals from KO1 ancestry. (B) Proportion of ancestry derived from central European southeast Iberia from the 3rd to 16th centuries CE (tables S20 and S21). Each bar Beaker/Bronze Age populations in Iberians from the Middle Neolithic to the represents one individual, with associated mtDNA (top) and Y-chromosome Iron Age (table S15). Colors indicate the Y-chromosome haplogroup for each male (bottom). Haplogroups with a likely recent nonlocal origin are bold.

Here we show that in Iberia, the allele continued 4. Q. Fu et al., Nature 534, 200–205 (2016). 17. T. Günther et al., Proc. Natl. Acad. Sci. U.S.A. 112, 11917–11922 to occur at low frequency in the Iron Age (fig. S9) 5. G. González-Fortes et al., Curr. Biol. 27,1801–1810.e10 (2017). (2015). and only approached present-day frequencies in 6. R. Martiniano et al., PLOS Genet. 13, e1006852 (2017). 18. G. Ruiz Zapatero, in Iberia. Protohistory of the Far West of 7. C. Valdiosera et al., Proc. Natl. Acad. Sci. U.S.A. 115, Europe: From Neolithic to Roman Conquest, the past 2000 years, pointing to recent strong 3428–3433 (2018). M. Almagro-Gorbea, Ed. (Universidad de Burgos, Fundación selection. 8. R. Fregel et al., Proc. Natl. Acad. Sci. U.S.A. 115, 6774–6779 Atapuerca, 2014), pp. 195–215. Beyond the specific insights about Iberia, this (2018). 19. M. Almagro-Basch, Ampurias. Historia de la ciudad y guía de las study serves as a model for how a high-resolution 9. I. Olalde et al., Nature 555, 190–196 (2018). excavaciones (Instituto Español de Prehistoria del CSIC y – Servicio de Investigaciones Arqueológicas de la Diputación ancient DNA transect continuing into historical 10. I. Lazaridis et al., Nature 513, 409 413 (2014). 11. N. Patterson et al., Genetics 192, 1065–1093 (2012). Provincial, 1951). periods can be used to provide a detailed descrip- 12. W. Haak et al., Nature 522, 207–211 (2015). 20. K. R. Veeramah et al., Proc. Natl. Acad. Sci. U.S.A. 115, tion of the formation of present-day populations 13. M. Lipson et al., Nature 551, 368–372 (2017). 3494–3499 (2018). (Fig. 1E); future application of similar strategies 14. C. Blasco, C. Liesau, G. Delibes de Castro, E. Baquedano, 21. J. S. Gerber, The Jews of Spain: A History of the Sephardic will provide equally valuable insights in other M. Rodriguez, in El campaniforme en la Península Ibérica y su Experience (The Free Press, 1992). contexto europeo, M. Rojo, R. Garrido, I. García, Eds. 22. L. de Ligt, L. E. Tacoma, Eds., Migration and Mobility in the world regions. (Universidad de Valladolid-Junta de Castilla y León, 2005), Early Roman Empire (Brill, 2016). pp. 457–479. 23. M. R. Bierling, S. Gitin, The Phoenicians in Spain: An REFERENCES AND NOTES 15. M. van de Loosdrecht et al., Science 360, 548–552 (2018). Archaeological Review of the Eighth–Sixth Centuries B.C.E. 1. N. Rohland, E. Harney, S. Mallick, S. Nordenfelt, D. Reich, 16. C. Liesau, E. Moreno, in Marfil y elefantes en la Península (Eisenbrauns, 2002). Philos. Trans. R. Soc. London Ser. B 370, 20130624 (2015). Ibérica y el Mediterráneo occidental, A. Banerjee, J. A. López 24. P. A. Zalloua et al., Am.J.Hum.Genet.83,633–642 2. I. Mathieson et al., Nature 528, 499–503 (2015). Padilla, T. X. Schuhmacher, Eds., Actas del coloquio (2008). 3. Q. Fu et al., Nature 524, 216–219 (2015). internacional (2012), pp. 87–98. 25. C. Bycroft et al., Nat. Commun. 10, 551 (2019).

Olalde et al., Science 363, 1230–1234 (2019) 15 March 2019 4of5 RESEARCH | REPORT

26. T. Bersaglieri et al., Am. J. Hum. Genet. 74, 1111–1120 by IT622-13 Gobierno Vasco, Diputación Foral de Álava, and V.S.G., E.G.-D., A.M.H.-C., J.J.-C., C.L., F.J.L.-C., D.L.-R., S.B.M., (2004). Diputación Foral de Gipuzkoa. We acknowledge support from M.M.P., A.O.F., G.P.B., P.R., M.S.B., A.C.S., J.M.V.E., M.Si., M.B.R., the Portuguese Foundation for Science and Technology K.W.A., W.H., R.P., C.L.-F., and D.R. assembled archaeological ACKNOWLEDGMENTS (PTDC/EPH-ARQ/4164/2014) and the FEDER-COMPETE 2020 material. I.O., S.M., N.P., M.F.-B., V.V.-M., M.Si., C.J.E., F.G., M.P., We thank I. Mathieson, M. Lipson, I. Lazaridis, J. Sedig, and K. Sirak project 016899. P.S. was supported by the FCT Investigator Program P.S., and D.R. analyzed data. I.O., C.L.-F., and D.R. wrote the for discussions, and M. E. Allentoft, K.-G. Sjögren, K. Kristiansen, (IF/01641/2013), FCT IP, and ERDF (COMPETE2020 – POCI). manuscript. Competing interests: The authors declare no and E. Willerslev for facilitating sample collection. We thank M.Si. and K.D. were supported by a Leverhulme Trust Doctoral competing interests. Data and materials availability: Sequencing M. Meyer for sharing the optimized oligo sequences for Scholarship awarded to M.B.R. and M.P. D.R. was supported by data are available from the European Nucleotide Archive, single-stranded library preparation. We thank the different museums an Allen Discovery Center grant from the Paul Allen Foundation, accession PRJEB30874; genotype dataset is available as (listed in the supplementary materials) for permission to study NIH grant GM100233, and the Howard Hughes Medical Institute. supplementary material. archaeological remains. Funding: J.M.F., F.J.L.-C., J.I.M., F.X.O., J.D., V.V.-M. and W.H. were supported by the Max Planck Society. and M.S.B. were supported by HAR2017-86509-P, HAR2017-87695-P, Authors contributions: N.R., N.A., N.B., O.C., B.J.C., D.F., A.M.L., SUPPLEMENTARY MATERIALS and SGR2017-11 from the Generalitat de Catalunya, AGAUR M.M., J.O., K.S., Z.Z., M.Si., K.D., C.J.E., D.J.K., M.B.R., W.H., R.P., www.sciencemag.org/content/363/6432/1230/suppl/DC1 agency. C.L.-F. was supported by Obra Social La Caixa and by and D.R. performed or supervised laboratory work. J.M.J.A., I.J.T.M., Supplementary Text FEDER-MINECO (BFU2015- 64699-P). L.B.d.L.E. was supported by D.C.S.-G., P.C., M.Sa., J.T., M.L., J.F.-E., J.A.M.-A., C.Ba., F.J.B., Figs. S1 to S11 REDISCO-HAR2017-88035-P (Plan Nacional I+D+I, MINECO). C.L., J.B., N.C., E.V.M., D.V., A.C., J.M.F., O.G.-P., J.I.M., F.X.O., J.M.V., Tables S1 to S22 P.R., and C.Bl. were supported by MINECO (HAR2016-77600-P). A.D.-C., I.O.-C., P.G.B., A.M.S., C.A.-F., J.J.E., A.M.-M., P.R.-G., J.R.M., References (27–189) A.Esp., J.V.-V., G.D., and D.C.S.-G. were supported by MINECO E.V.V., K.T.L., J.M., A.W., G.D., B.A., F.C., A.Esp., G.d.P., A.Est., Genotype Dataset (HAR2009-10105 and HAR2013-43851-P). D.J.K. and B.J.C. were C.F., G.F., S.F., F.G.-G., T.M., A.R., J.V.-V., G.A.A., V.B.G., L.B.d.L.E., supported by NSF BCS-1460367. K.T.L., A.W., and J.M. were M.B.S., G.G.A., M.S.H.P., A.L., Y.C.M., I.C.B., A.F.F., D.L.-S., M.S.T., 12 September 2018; accepted 30 January 2019 supported by NSF BCS-1153568. J.F.-E. and J.A.M.-A. were supported A.C.V., C.Bl., J.D., M.J.d.P.M., A.A.D.-C., R.F.F., J.F.F., R.G.-P., 10.1126/science.aav4040 Downloaded from http://science.sciencemag.org/

on March 14, 2019

Olalde et al., Science 363, 1230–1234 (2019) 15 March 2019 5of5 The genomic history of the Iberian Peninsula over the past 8000 years Iñigo Olalde, Swapan Mallick, Nick Patterson, Nadin Rohland, Vanessa Villalba-Mouco, Marina Silva, Katharina Dulias, Ceiridwen J. Edwards, Francesca Gandini, Maria Pala, Pedro Soares, Manuel Ferrando-Bernal, Nicole Adamski, Nasreen Broomandkhoshbacht, Olivia Cheronet, Brendan J. Culleton, Daniel Fernandes, Ann Marie Lawson, Matthew Mah, Jonas Oppenheimer, Kristin Stewardson, Zhao Zhang, Juan Manuel Jiménez Arenas, Isidro Jorge Toro Moyano, Domingo C. Salazar-García, Pere Castanyer, Marta Santos, Joaquim Tremoleda, Marina Lozano, Pablo García Borja, Javier Fernández-Eraso, José Antonio Mujika-Alustiza, Cecilio Barroso, Francisco J. Bermúdez, Enrique Viguera Mínguez, Josep Burch, Neus Coromina, David Vivó, Artur Cebrià, Josep Maria Fullola, Oreto García-Puchol, Juan Ignacio Morales, F. Xavier Oms, Tona Majó, Josep Maria Vergès, Antònia Díaz-Carvajal, Imma Ollich-Castanyer, F. Javier López-Cachero, Ana Maria Silva, Carmen Alonso-Fernández, Germán Delibes de Castro, Javier Jiménez Echevarría, Adolfo Moreno-Márquez, Guillermo Pascual Berlanga, Pablo Ramos-García, José Ramos-Muñoz, Eduardo Vijande Vila, Gustau Aguilella Arzo, Ángel Esparza Arroyo, Katina T. Lillios, Jennifer Mack, Javier Velasco-Vázquez, Anna Waterman, Luis Benítez de Lugo Enrich, María Benito Sánchez, Bibiana Agustí, Ferran Codina, Gabriel de Prado, Almudena Estalrrich, Álvaro Fernández Flores, Clive Finlayson, Geraldine Finlayson, Stewart Finlayson, Francisco Giles-Guzmán, Antonio Rosas, Virginia Barciela González, Gabriel García Atiénzar, Mauro S. Hernández Pérez, Armando Llanos, Yolanda Carrión Marco, Isabel Collado Beneyto, David

López-Serrano, Mario Sanz Tormo, António C. Valera, Concepción Blasco, Corina Liesau, Patricia Ríos, Joan Daura, María Downloaded from Jesús de Pedro Michó, Agustín A. Diez-Castillo, Raúl Flores Fernández, Joan Francès Farré, Rafael Garrido-Pena, Victor S. Gonçalves, Elisa Guerra-Doce, Ana Mercedes Herrero-Corral, Joaquim Juan-Cabanilles, Daniel López-Reyes, Sarah B. McClure, Marta Merino Pérez, Arturo Oliver Foix, Montserrat Sanz Borràs, Ana Catarina Sousa, Julio Manuel Vidal Encinas, Douglas J. Kennett, Martin B. Richards, Kurt Werner Alt, Wolfgang Haak, Ron Pinhasi, Carles Lalueza-Fox and David Reich http://science.sciencemag.org/ Science 363 (6432), 1230-1234. DOI: 10.1126/science.aav4040

Genomics of the Iberian Peninsula Ancient DNA studies have begun to help us understand the genetic history and movements of people across the globe. Focusing on the Iberian Peninsula, Olalde et al. report genome-wide data from 271 ancient individuals from Iberia (see the Perspective by Vander Linden). The findings provide a comprehensive genetic time transect of the region. Linguistics analysis and genetic analysis of archaeological human remains dating from about 7000 years ago to the present elucidate the genetic impact of prehistoric and historic migrations from Europe and North Africa. Science, this issue p. 1230; see also p. 1153

on March 14, 2019

ARTICLE TOOLS http://science.sciencemag.org/content/363/6432/1230

SUPPLEMENTARY http://science.sciencemag.org/content/suppl/2019/03/13/363.6432.1230.DC1 MATERIALS

RELATED http://science.sciencemag.org/content/sci/363/6432/1153.full CONTENT

Use of this article is subject to the Terms of Service

Science (print ISSN 0036-8075; online ISSN 1095-9203) is published by the American Association for the Advancement of Science, 1200 New York Avenue NW, Washington, DC 20005. 2017 © The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. The title Science is a registered trademark of AAAS. REFERENCES This article cites 144 articles, 14 of which you can access for free http://science.sciencemag.org/content/363/6432/1230#BIBL

PERMISSIONS http://www.sciencemag.org/help/reprints-and-permissions Downloaded from http://science.sciencemag.org/

on March 14, 2019

Use of this article is subject to the Terms of Service

Science (print ISSN 0036-8075; online ISSN 1095-9203) is published by the American Association for the Advancement of Science, 1200 New York Avenue NW, Washington, DC 20005. 2017 © The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. The title Science is a registered trademark of AAAS.